ISSN 0373-580 X Bol. Soc. Argent. Bot. 40 (3-4): 241 - 281. 2005
Epidermal characteristics of toxic plants for cattle from the Salado river basin (Buenos Aires, Argentina)
SUSANA E. FREIRE1,2, ANA M. ARAMBARRI1, NÉSTOR D. BAYÓN1, GISELA SANCHO1,2, ESTRELLA URTUBEY2, CLAUDIA MONTI1, MARÍA C. NOVOA1, and MARTA N. COLARES1
Summary: One hundred and eighty species belonging to 41 families inhabiting the Salado River Basin of the province of Buenos Aires (Argentina) were previously reported to be toxic for cattle. The purpose of this study was to provide a tool to distinguish the taxa when the plant material is desintegrated. In this way, an approach to the identification of these taxa through leaf epidermal features (anticlinal epidermal cell wall patterns, cuticular ornamentation, stomata, and hair types) is performed. A key to the 180 species as well as illustrations of diagnostic characters are given.
Key words: Buenos Aires, Salado River Basin, toxic plants, anatomy, epidermal characters, stomata, hairs, Dicotyledons, Monocotyledons.
Resumen: Caracteres epidérmicos de las plantas tóxicas para el ganado de la Depresión del Salado (Buenos Aires, Argentina). Las plantas tóxicas para el ganado están representadas en la Depresión del Salado (provincia de Buenos Aires, Argentina) por 180 especies pertenecien- tes a 41 familias. El objetivo del presente trabajo es determinar estos taxa a partir de material desintegrado, utilizando caracteres epidérmicos foliares (paredes anticlinales de las células epidérmicas, ornamentación de la cutícula, tipos de estomas y pelos). Se brinda una clave para la determinación de las especies e ilustraciones de los caracteres diagnósticos.
Palabras clave: Buenos Aires, Depresión del Salado, anatomía, caracteres epidérmicos, estomas, pelos, Dicotiledóneas, Monocotiledóneas, plantas tóxicas.
Introduction The political province of Buenos Aires is situated in central eastern Argentina. It is covered in most of its surface by a herbaceous grassy steppe, called “pampas.” The species studied are from a plain with a poorly developed drainage system. This area is known as Salado River Basin which represents approximately 80,000 km2 (Fig. 1). The principal economic activity in this area is the cattle breeding, based on natural pastures. The knowledge of the vegetation of this area is relevant for human development. Numerous floristic studies have been carried out in this area (Cabrera, 1963-1967; Vervoorst, 1967; Cabrera & Zardini, 1978; Cabrera et al., 2000). Within the diverse plant families inhabiting the Sa- lado River Basin, 180 species belonging to 41 families show toxicity for cattle (Casós, 1935; Rate- ra, 1945; Tokarnia & Dobereiner, 1982; Ragonese & Milano, 1984; Gallo, 1979; Pertusi, 1987). Epidermal traits, i.e. epicuticular wax deposition, cuticular 1 Área de Botánica, Departamento de Ciencias Biológi- cas, Facultad de Ciencias Agrarias y Forestales, UNLP, calle 60 y 119, C.C. 31, 1900 La Plata, Argentina). 2 División Plantas Vasculares, Museo de La Plata, Paseo Fig. 1. Map of Buenos Aires province showing the Salado del Bosque s/n, 1900 La Plata, Argentina. River Basin (area in gray).
241 Bol. Soc. Argent. Bot. 40 (3-4) 2005 ornamentation, epidermal cells, stomata, and hairs, have patterns used in (Table 1) was adapted from Stace proved to be an important tool in taxa delimitation in (1965): Stace’s types 1 and 2 correspond to type 1 many plant families (Metcalfe & Chalk, 1950-1979; here; Stace’s types 3 and 4 correspond to type 2 Uphof et al., 1962; Sinclair & Sharma, 1971; Lackey, here; Stace’s types 5 and 6 correspond to type 3 1978; Arambarri & Colares, 1993; Ditsch et al., 1995; here; Stace’s types 7 and 8 correspond to type 4 Barthlott et al., 1998; Stenglein et al., 2003) and also here. The anticlinal epidermal cell wall patterns are in distinguishing fragmented vegetables from feces indicated in Table 1 as: adaxial surface / abaxial and stomach contents being resistant to the digestive surface. process (Yagueddú & Cid, 1992; Pelliza et al., 1997; Stomata types were classified according Cid & Sierra, 2004). Therefore, it would be interesting to seek morphological traits that led to diagnose on causes Metcalfe & Chalk (1950, 1979) and Van Cotthem of animal mortality through feces and stomach contents. (1970), however, to establish the monocotyledons In order to achieve this goal, we apply herein histo- types, we followed Fryns-Claessens & Van morphological characters, such as leaf epidermal features. Cotthem’s (1973) classification. Leaf margin (visible in transparent leaves) was Material and Methods only used to distinguish two groups of species with glabrous leaves. Plant material studied The nomenclature follows Zuloaga et al.(1994) The study was performed using fresh leaves and Zuloaga & Morrone (1996, 1999). (collected in Buenos Aires province), and dried Epidermal characters in Poaceae were described leaves taken from herbarium specimens belonging according to the terminology of Metcalfe (1960) and to LP, LPAG, LPS, and SI (acronyms according Ellis (1979), and hair terminology follows Metcalfe Holmgren et al., 1990). The 180 taxa investigated & Chalk (1950, 1979) and Uphof (1962). For glandu- and vouchers are detailed in Appendix 1. lar hairs (Table 1) the number in brackets indicates Methods the number of head cells, and for non-glandular Fully expanded leaves were selected for the hairs the number of hair cells above the epidermis. study. Data were obtained from the central area of The observed hairs and stomata types are described the midlamina on both surfaces. For reconstitution according to Metcalfe & Chalk (1979), Ramayya of dried leaves we followed D’Ambrogio de (1962), and Uphof (1962). Argüeso (1986). Then, the material was fixed in formalin, glacial acetic acid, and 50% ethanol at a Results and Discussion 5:5:90 ratio (F.A.A.). Most of the epidermal From the 41 studied families inhabiting the Sala- microcharacters were studied by peeling and/or in do River Basin, the most representated were: samples cleared using the technique of Dizeo de Asteraceae (41 spp.), Poaceae (16 spp.), Solanaceae Strittmatter (1973). However, the replica method (14 spp.), Fabaceae (13 spp.), and Brassicaceae (10 (according to Freeman, 1984) was used in some taxa spp.). The results are presented in Table 1. where it was not possible to get epidermis by Even if stomata and indumentum types of the peeling or chloral hydrate clearing. To study the species studied are basically in agreement with those epidermal characters of the species belonging to described by Metcalfe (1960), Metcalfe & Chalk (1979, Poaceae we followed the technique of Metcalfe (1960). 1989) and Uphof (1962) for all families studied, a more The semipermanent slides were stained using saffranin detailed discussion of some families is included, in 80% ethanol and mounted in gelatine-glycerine. because they have some microcharacters that would Observations, and original drawings were made be cited for the first time and/or they present some with a light microscope, Leitz SM lux with camera luci- special traits. They are as follows: da. Measurements of stomata (length and width) and hairs were taken using a Nikon light microscope Apiaceae (6 spp. surveyed): We found equipped with an ocular micrometer. The average size anomocytic, diacytic, and paracytic stomata types; of hairs and stomata were determined based on all species showed glabrous epidermal surfaces, measurements performed on 15-20 replicates per sample. although papillae were found at midvein level in Cuticular ornamentation was cited only when it two species of Ammi. It is the first time in which was conspicuous (Table 1) diacytic stomata type has been cited for Apiaceae, The classification of anticlinal epidermal cell wall whereas anomocytic and paracytic have been
242 S. E. Freire et al., Epidermal characteristics of toxic plants for cattle Table 1. Epidermal characters of the toxic plants of Salado River Basin. Numbers in brackets by the species refer to bibliography cited below the table.
ab a th t b at
R h a ab f a Cy t T ch a t C ta a a a Cy acty Cy h t D ta a a a ch ch ac a ca y b v a tat a t c a c a at L t ch ach f L t f L L t t S h ha
A a a th hyb A v A a A v a a C ac at Cyc t hy y a c at c v a V ca a
243 Bol. Soc. Argent. Bot. 40 (3-4) 2005 Table 1. Continuation
A c a c a av ca A a ab achy t ha a ata O y ta a Achy c at A b at f a A th c t a A ct A t q a at q a at Baccha a t B a t c ata B c f a B t a B f c B t a B a a Ca aca th C ta aca c t a a C t C t t a C ch tyb C v a C y ab a b a Cy a aca c Ga a a a ta ca a ta ca y y a th cab a Lact ca a O aca th S c b a S c b a t a t t S bach bach S a a a ca S tw
244 S. E. Freire et al., Epidermal characteristics of toxic plants for cattle
Table 1. Continuation
S v a S yb a a S a ch ch S ch ac Ta t ta Ta a ac ff c a
V b a c V a b ca b ca W a a ca a th cava a th A ch a ff c a ch a ta
t a ca
B a ca a B a a Ca ab a a t Ca a a aba Ra ha a ha t R at v Ra t R a a t t aq at c S y b a t S O t aa chava ta a a tt
W at h aca tha
Sa b c a t a
A t a tha yca t t a hy Sa a a ff c a S aa v Ba a c a a
245 Bol. Soc. Argent. Bot. 40 (3-4) 2005 Table 1. Continuation
Ch a b C h c h c C a b atw Sa aka ka
C v v a v h b a athy a h t aha R c c A ab c Ga a ff c a L t c c at L ab L b t a t a b c ff c a a k aac ata S ac y b a T f V c a a a a a V at va a a aca ata ca ata ff c a
a v f a
c c ta a ac G a La a ca a b v a a va y v t
246 S. E. Freire et al., Epidermal characteristics of toxic plants for cattle
Table 1. Continuation
S a h b f a b c a t a y h ca B hav a ff a ca a ab a a a O a c c ata c c ata
a f aca a
hyt acca a ca a ca t t a a a ta a t a a t a L b a b a y av c a
c v v a ath f ca a R c R bt f R ch t aca ac a
A a a a v
A ca ta a ca ta a C at b a C t v Ra c a f
R cy ba a a
R cat R
247 Bol. Soc. Argent. Bot. 40 (3-4) 2005 Table 1. Continuation
D ca aa ca a B av t c ata Ga cha a cha a a a ahy a D av c a V ba c tha C t a q Dat af c t a a a ca
f a hy a v c a Sa ch a a f a S a b a S chac chac S c c S f S a a f S a c hy S y a y a S b bat T a at f a a ta a ff c a t ca
1) Metcalfe (1960); (2) Prat & Vignal (1968); (3) Caro & Sánchez (1969); (4) Sánchez (1971); (5) Gould & Shaw (1992); (6) Sánchez (1974); (7) Westerkamp & Demmelmeyer (1997); (8) Bayón & Arambarri (1999); (9) Selvi & Bigazzi (2001); (10) Di Fulvio (1976); (11) Amat (1988); (12) Gattuso & Gatusso (1998); (13) Ariza Espinar (1973); (14) Pertusi (1987); (15) Barboza et al. (2001); (16) Simón et al. (2002); (17) Cabrera (1944); (18) Arroyo (1986); (19) Yagueddú & Cid (1992); (20) Arambarri & Colares (1993); (21) Gatusso & Gatusso (1989); (22) Gatusso (1996); (23) Ragonese & Covas (1947); (24) Gatusso (2000); (25) Medán (1986); (26) Metcalfe & Chalk (1950); (27) Colares et al. (1999); (28) Cabrera (1979); (29) Stenglein (2001); (30) Bruno et al. (1999). H = hooks; P = prickles; Pa = papillae
248 S. E. Freire et al., Epidermal characteristics of toxic plants for cattle
Fig. 2. Epidermal characters of Poaceae. a, stoma with parallel-sided subsidiary cells: Briza minor; b, stoma with triangular subsidiary cells: Cynodon dactylon; c, stoma with dome subsidiary cells: Leptochloa chloridiformis; d, micro-hair with almost hemispherical distal cell: Eragrostis cilianensis; e, dumb-bell silica-body: Echinochloa crusgalli; f, saddle-shaped silica-body: Eragrostis cilianensis; g, rectangular silica-body: Lolium multiflorum; h, cross-like silica-body: Sorghum halepense; i, micro-hairs with pear-like distal cell: Eleusine indica; j, micro-hair with rod-like distal cell: Digitaria sanguinalis. Scale bar = 50 µm. previously mentioned by Metcalfe & Chalk (1979). nal and basal cells (similar to bristle hair, according Types of stomata varied notably in such a small to the illustration); whip hairs (i.e. aseptate flagellate group of genera and species. This variability was according to Ramayya, 1962); two-armed (i.e. T- previously noticed by Forcone & Ayestaran (1996). shaped); stellate; candelabra hairs; shaggy hairs, Eryngium epidermal cells showed parallel and peltate scales. Uniseriate like bristles, peltate distribution and also paracytic stomata, which was scales, and candelabra hairs, were mentioned by noted before by Yagueddú & Cid (1992). these authors for genera not included in the present study, whereas stellate or branched hairs cited by Asclepiadaceae (5 spp. analyzed): Results agree them for the genus Baccharis were found in other with Metcalfe & Chalk (1950, 1979) and also with species not included in this paper (Freire, in prep.). Bayón & Arambarri (1999) who mentioned that All other hair types, i.e. whip hairs, two-armed or paracytic was the most frequent type of stomata. T-shaped, and shaggy hairs, agree with those Bayón & Arambarri (l.c.), also illustrated the described in this study. In addition, we found epidermal and trichome ornamentation, specially for conical hairs in nine species, in two of which Oxypetalum solanoides which present hairs, (Ambrosia tenuifolia, Aster squamatus), they were exclusively waxes ornamentated on the apical cell. previously reported by Yagueddú & Cid (1992), whereas Barboza et al. (2001) illustrated conical Asteraceae (41 spp. analyzed): Metcalfe & Chalk hairs in one species of Xanthium (X. spinosum). In (1950, 1979) characterized Asteraceae by having the agreement with Metcalfe & Chalk (1950, 1979), glan- following seven hairs: uniseriate consisting of dular hairs are widely distributed (only few species uniform cells apart from modification of the termi- of the total species analyzed have not glandular
249 Bol. Soc. Argent. Bot. 40 (3-4) 2005
Fig. 3. Hairs of Asclepiadaceae, Boraginaceae, Brassicaceae, Cactaceae, Caprifoliaceae, Caryophyllaceae, Chenopodiaceae,
Convolvulaceae, Geraniaceae, and Lamiaceae. a, conical: Morrenia odorata; b, conical: Oxypetalum solanoides; c, falcate, c1, glandular capitate: Anchusa officinalis; d, conical: Echium plantagineum; e, conical, e1, glandular capitate: Heliotropium amplexicaule; f, moniliform: Parodia ottonis; g, short bristle: Sambucus australis; h, bristle: Agrostemma githago; i, moniliform: Saponaria officinalis; j, vesicular: Chenopodium album; k, whip: Ch. murale; l, conical: Bassia scoparia; m, conical: Salsola kali; n, glandular capitate, n1, conical: Convolvulus arvensis; o, conical: Brassica nigra; p, 3-armed: Capsella bursa-pastoris; q, falcate: Cardaria draba; r, barrel-shaped: Raphanus sativus; s, falcate: Rapistrum rugosum; t, long-conical:
Sisymbrium altissimum; u, glandular capitate bottle-shaped: u1, bristle: Erodium malacoides; x, conical: Geranium molle; y, conical, y1, glandular capitate: Lamium amplexicaule; z, porrect stellate, z1, glandular capitate: Marrubium vulgare. Scale bars: a, b, c1, g, j-o, q, z = 50 µm; c, d = 300 µm; e, e1, h, i, p, r, s, x, y = 100 µm; f, t = 200 µm; y1,z1=25µm.
250 S. E. Freire et al., Epidermal characteristics of toxic plants for cattle
Fig. 4. Hairs of Asteraceae. a, whip: Centaurea calcitrapa; b, whip: Carduus acanthoides; c, whip and d, shaggy: Cirsium vulgare; e, conical: Conyza bonariensis; f, conical: Gaillardia megapotamica; g, whip: Senecio grisebachii; h, conical: Solidago chilensis; i, conical: Verbesina encelioides; j, conical: Wedelia glauca; k, whip: Onopordon acanthium; l, conical: Xanthium cavanillesii; m, conical: Xanthium spinosum; n, whip: Achyrocline satureioides; o, conical: Ambrosia tenuifolia; p, whip: Arctium minus; q, T-shaped: Anthemis cotula; r, whip: Aster squamatus; s, whip: Baccharis artemisoides; t, pilose nest: B. rufescens; u, pilose nest: B. notosergila; v, conical: Bidens pilosa. Scale bars: a-e, g-u = 50 µm; v, f = 100 µm.
251 Bol. Soc. Argent. Bot. 40 (3-4) 2005
Fig. 5. Hairs of Fabaceae, Malvaceae, Martyniaceae, Nyctaginaceae, Oxalidaceae, Plantaginaceae, Polygonaceae, Primulaceae, Ranunculaceae, Rhamnaceae, and Rubiaceae. a, conical: Adesmia bicolor; b, bristle: Galega officinalis; c, bristle: Lotus corniculatus; d, conical: Lupinus gibertianus; e, bristle, e1, glandular capitate: Melilotus indicus; e2, glandular capitate: M. albus; f, bristle: Parkinsonia aculeata; g, bristle: Senna corymbosa; h, glandular uniseriate: Trifolium repens; i, bristle, i1, glandular capitate: Vicia graminea; j, conical, j1-j2, glandular capitate: Malva sylvestris; k, stellate multiangulate, k1-k2, glandular capitate: Sida rhombifolia; l, glandular capitate with head cells radiate, Ibicella lutea; m, glandular capitate:
Boerhavia diffusa var. leiocarpa; m1, glandular capitate: Mirabilis jalapa; n, bristle: Oxalis corniculata; o, glandular capitate: Plantago australis; p, shaggy, p1, conical-flagellate, p2, glandular capitate: Polygonum lapathifolium; q, short conical: P. convolvulus; r, glandular capitate: Anagallis arvensis; s, bristle, s1, glandular clavate: Anemone decapetala; t, glandular clavate: Clematis bonariensis; u, bristle: C. montevidensis; x, bristle: Ranunculus repens; y, bristle: Discaria americana;z, barrel shaped: Borreria verticillata. Scale bars: a-i, n, p, p1,q,r,s,u,x,z=100µm;j,m1, o, s1,t,y=50µm;p2=25µm.
252 S. E. Freire et al., Epidermal characteristics of toxic plants for cattle
Fig. 6. Hairs of Sapindaceae, Scrophulariaceae, Solanaceae, Turneraceae, and Urticaceae. a, peltate scale, a1, bristle: Dodonaea viscosa; b, stellate stalked: Verbascum thapsus; c, glandular capitate: d, conical: Cestrum parqui; e, conical, f, glandular capitate: Datura ferox; g, conical: Nicotiana longiflora; h, i, glandular capitate, j, conical: Salpichroa origanifolia; k, dendritic: Physalis viscosa; l, conical: Solanum chacoense; m, stellate stalked: S. elaeagnifolium; n, Y-shaped: S. diflorum; o, bristle, p, glandular shaggy: Turnera sidoides subsp. pinnatifida; q, conical: Parietaria officinalis; r, bristle, r1, stinging:
Urtica urens. Scale bars: a, a1,b,m,n,p,r=50µm;c,d,h-k=200µm;e,f,l,o,q,r1= 100 µm; g = 300 µm.
253 Bol. Soc. Argent. Bot. 40 (3-4) 2005
Fig. 7. Stomatal types. a-f, anysocytic: a, Polygonum aviculare; b, Raphanus sativus; c, Sida rhombifolia; d, Turnera sidoides subsp. pinnatifida; e, Adesmia bicolor; f, Nicotiana glauca; g, polycitic: Plantago australis; h, cyclocytic: Baccharis notosergila; i, parallelocytic: Portulaca oleracea; j, k, diacytic: j, Lamium amplexicaule; k, Ammi majus; l, hexacytic: Opuntia arechavaletae; m-r, paracytic: m, Convolvulus arvensis; n, Oxypetalum solanoides; o, Galium richardianum; p, Ricinus communis; q, Senna corymbosa;r,Vinca major. Scale bars: a-d, g, j-p,r=50µm;e,f,h,i,q=100µm.
254 S. E. Freire et al., Epidermal characteristics of toxic plants for cattle
Fig. 8. Epidermal special characters. a, vesicular cells: Polygonum persicaria; b, hydropoten: Nymphoides indica; c, cystholits: Urtica urens; d, “Licópoli glands”: Limonium brasiliense; e, thick anticlinal walls: Wigginsia tephracantha; f, cuticular ornamentation, striate: Ranunculus cymbalaria. Scale bars: 100 µm.
255 Bol. Soc. Argent. Bot. 40 (3-4) 2005 hairs), and they are ususally 2-seriate, occasionally presented ornamented 1-celled hairs only as an depressed below the leaf surface (i.e. Hymenoxys exception, i.e. Brassica rapa and Sisymbrium irio. anthemoides). In many species of Baccharis glan- dular and non glandular hairs appear forming tufts Cactaceae (3 spp. studied): This family is or pilose nest (Ariza Espinar, 1973; Pertusi, 1987; distinguished by hexacytic and parallelocytic stomata, Helwig, 1992). Four species of the six studied have and moniliform hairs. Opuntia presented hexacytic pilose nest, the remaining two, have isolated non- stomata and glabrous surfaces whereas Parodia and glandular hairs. Stomata are predominantly Wigginsia exhibited parallelocytic stomata and anomocytic, however, three species of Baccharis moniliform hairs. These results and the presence of show ciclocytic stomata. This stomata type was not thick anticlinal epidermal cell walls in Wigginsia are mentioned neither by Metcalfe & Chalk (1950, 1979), wholy in agreement with characters reported by Di for Asteraceae nor by Ariza Espinar (1973) for Fulvio (1976) and Metcalfe & Chalk (1979). Baccharis. However, ciclocytic stomata were reported for other genera of Asteraceae (Freire, Caprifoliaceae: The only studied species, 1986; Crisci & Freire, 1986; Anderberg & Freire, Sambucus australis, showed anomocytic stomata 1991; Freire, 1993) and for Baccharis (Pertusi, 1987). and non-glandular hairs. Both features correspond to data reported by Metcalfe & Chalk (1979) and Boraginaceae (3 spp. analyzed): All the species also with those cited for Sambucus nigra by showed anomocytic stomata and two types of hairs: Ponessa & Parrado (2001). (1) simple, unicellular, conical or falcate; (2) capitate with unicellular head. Those features correspond Caryophyllaceae (4 spp. studied): We found two well with Metcalfe & Chalk (1950, 1979) who named stomata types: diacytic and anomocytic. Metcalfe & the former type (i.e., unicellular conical or falcate), Chalk (1950, 1979) mentioned anisocytic ones but we as boraginaceous. Patel & Inmandar (1971) studied did not observe this third type. We saw that three of 10 species of Boraginaceae and found that eight of the four species have non-glandular hairs, the third them presented three types of stomata: anomocytic (Spergula arvensis) also has glandular capitate hairs (ranunculaceous type of Vesque, 1889), paracytic with an unique cell in the apex in according to the (rubiaceous type of Vesque, 1889), and anisocytic results of Metcalfe & Chalk (1979). Furthermore, we (cruciferous type of Vesque, 1889), although observed bristles hairs in Agrostemma githago a anomocytic was present in all genera. According character that was not mentioned for Caryophyllaceae to these authors, Heliotropium presented also by Metcalfe & Chalk (1979). diacytic type (caryophyllaceous type of Vesque, 1889). We found, in accordance with Selvi & Bigazzi Chenopodiaceae (6 spp. analyzed): Five species (2001), anomocytic stomata type, and from seven showed anomocytic stomata whereas one (Salsola) hair types mentioned by them we found only hair presented paracytic ones. These results are types 1 and 3 in Anchusa, Echium, and consistent with those exposed by Metcalfe & Chalk Heliotropium. These results corroborated the (1950, 1979). Barboza et al. (2001) found in epidermal traits mentioned by Barboza et al. (2001) Chenopodium species anomocytic, anisocytic, and and Monti et al. (2003). tetracytic stomata types, however they also mentioned as the most frequent the anomocytic Brassicaceae (10 spp. examined): They type. According to our observations there are presented two stomata types: anomocytic and different types of hairs: 1) conical, 2) whip and, 3) anisocytic. Ancibor (1984) and Barboza et al. (2001) vesicular. A great variability of hairs was also found cited both types of stomata. Anisocytic type was by Barboza et al. (2001). We considered, as Metcalfe considered a diagnostic character for this family by & Chalk (1950) did, that Chenopodium and Salsola Metcalfe & Chalk (1979), and it has been found by have uniseriate thin-walled hairs and we can add Bassia Arroyo (1984, 1986) and Diorio (1986) in different to the list. In the toxic species of Chenopodium we species of this family. Different types of hairs were noticed vesicular hairs with stalks with one or more observed: (1) conical, (2) falcate, (3) stellate sessile, cells. They were named “salt glands” by Ancibor (4) 3-armed, and (5) barrel shaped. All these hair types (1992) and D’Ambrogio et al. (2000). There are three were considered frequent within the family by Metcalfe ruderal species of the genus Chenopodium known as & Chalk (1950). Although conical hairs are similar to “paicos” which are considered toxic for human those observed in Solanaceae, the family Brassicaceae consumption (Simon, 1987) but not for cattle, so we
256 S. E. Freire et al., Epidermal characteristics of toxic plants for cattle did not include them in our study. two hairs types: (1) bristles and (2) glandular capitate with 1-celled head. Our results are in Convolvulaceae: The only species analyzed of this agreement with Metcalfe & Chalk (1950). family presented paracytic stomata and conical 1-celled Furthermore, anomocytic stomata type is additioned hairs and glandular capitate hairs with a variable to the paracytic type reported by Metcalfe & Chalk number of cells. All characters are completely in (1979) for this family. coincidence with Metcalfe & Chalk (1950, 1979). The stomata type also agree with epidermal traits reported by Passifloraceae: Only Passiflora caerulea was Yagueddú & Cid (1992) for other species of this family. studied, which presented anomocytic stomata type as was mentioned by Metcalfe & Chalk (1950). The Euphorbiaceae (4 spp. analyzed): We found same authors (1979) included the paracytic type and anomocytic stomata in Euphorbia lathyrus and E. reported the presence of hairs as characteristic of peplus, and paracytic in Manihot grahamii and the family. Barboza et al. (2001) studied this species Ricinus communis, both stomata types were reporting in addition two types of stomata that we mentioned by Metcalfe & Chalk (1950) for this have not seen: anisocytic and paracytic. In family, but they reported anomocytic type as the according to that paper we did not find hairs. usual type. Our results agree with Metcalfe & Chalk (1979), however, we did not find anisocytic type Phytolaccaceae (3 spp. studied): They were mentioned by them. In spite of the fact that all the characterized by the presence of anomocytic species studied may be considered glabrous, stomata type and infrequent simple, uniseriate Ricinus communis presented a few short-conical 1- conical hairs. These epidermal features coincide with celled hairs sometimes appearing as papillae. Within Metcalfe & Chalk (1950, 1979) and Gattuso (1996). the group of species studied we did not find glan- According to Gattuso (1996) anisocytic and dular hairs which were mentioned by Metcalfe & paracytic stomata are rare in Phytolaccaceae. Chalk (1950, 1979) for Euphorbiaceae. All our results coincide with those reported by Barboza et al. (2001) Plantaginaceae: Only Plantago australis var. for Euphorbia serpens. australis was examined. It was characterized by having predominantly polocytic stomata, i.e. stomata Fabaceae (13 spp. examined): This family was attached to the distal side of the single subsidiary characterized by having bristles hairs, glandular hairs, cell, only a few ones are anomocytic. Polocytic and anisocytic and anomocytic stomata. The genus stomata was previously reported by Van Cotthem Senna was an exception, having paracytic stomata. (1970) in ferns. Metcalfe & Chalk (1979), Yagueddú Both epidermal traits are in agreement with Metcalfe & Cid (1992), and Barboza et al. (2001), reported & Chalk (1950, 1979) and numerous authors like anomocytic stomata in other species of Plantago. Ragonese (1969), Lackey (1978), Soladoye (1982), However, the illustrations of P. lanceolata by Yagueddú & Cid (1992), Ponesa et al. (1998), Barboza Barboza et al. (2001) and by Yageddú & Cid (1992) et al. (2001), Stenglein et al. (2003, 2004). reveal that although the stomata are predominantly anomocytic, a few ones can be considered as Menyanthaceae: Only Nymphoides indica was polocytic. Although Metcalfe & Chalk (1979) and examined. It was characterized by anomocytic Barboza et al. (2001) found different types of hairs stomata on the adaxial surface and the presence of within Plantago, we are able only to report the hydropoten on the abaxial epidermis. Both presence of glandular capitate hairs with 2-celled head. diagnostic features accord with Metcalfe & Chalk (1979) where the authors described extensively the Poaceae (16 spp. examined): Three characters characteristics of the hydropoten (water drinks) were mainly employed to distinguish these species: structure. Hydropoten on abaxial epidermis was shape of silica-bodies, shape of stomata subsidiary previously mentioned for this species by Gattuso cells, and micro-hairs. They are discussed below & Gattuso (1989). arranged by types of epidermis defined by Prat (1936), Parodi (1958), and Tateoka et al. (1959). Oxalidaceae: Oxalis corniculata var. Festucoid type: eigth of the 16 species showed corniculata was the only species studied. It was this type. No-one had micro-hairs. Four species characterized by having anomocytic stomata and (Briza minor, Elymus breviaristatus, Holcus
257 Bol. Soc. Argent. Bot. 40 (3-4) 2005 lanatus, and Hordeum murinum) presented stomata the group of glandular hairs a special type with parallel-sided subsidiary cells, one (Cortaderia described as superficial mucilage glands, usually selloana) stomata with low domed-shaped present in young leaves. Then, these authors (1979) subsidiary cells and three species of Lolium showed gave a detailed explanation of this special “hairs” both types of subsidiary cells. In this group, the named hydropoten and mentioned Polygonaceae shape of silica-bodies varied within this range of as one of the dicotyledoneous families in which shapes: quadrangular, rectangular, rounded, or they found them. Mitchell (1971) described the elongated. All these features are tightly in agreement derived structure of these hairs as “valvate with Metcalfe´s results (1960). chambers”. Some species also presented vesicular Chloridoid type: five species presented this type: cells, and these were mentioned by Gattuso (2000). Cynodon dactylon, C. hirsutus, Eleusine indica, Eragrostis cilianensis, and Leptochloa Rhamnaceae: Only Discaria americana was chloridiformis. They showed saddle-shaped silica- studied, which presented anomocytic stomata and bodies and micro-hairs with hemispherical apical cell. bristles. These characters are in agreement with Eleusine indica is an exception because of its Metcalfe & Chalk (1950, 1979) and Medan (1986). characteristic pear-like microhairs, which were studied by several authors like Metcalfe (1960), Prat Rutaceae: Fagara hyemalis, the only species & Vignal (1968), and Sánchez (1974). Metcalfe analyzed, showed anomocytic stomata and stellate considered that this micro-hairs can also be regarded hairs. Metcalfe & Chalk (1950) considered that this as sunken papillae or very small macro-hairs. family shows different stomata types whilst in 1979 Eleusine indica presented stomata with triangular they cited paracytic but not anomocytic stomata. subsidiary cells whereas the rest of species showed Barboza et al. (2001) also found anomocytic stomata both triangular or low domed-shaped ones. Other type in this family. Metcalfe & Chalk (1950, 1979) characters, such as macrohairs, prickles and papillae, also reported the presence of the stellate hairs. found here in genus Cynodon, were also reported by other authors (Prat & Vignal, 1968; Caro & Solanaceae (14 spp. surveyed): They had mainly Sánchez , 1969; Sánchez, 1971). two types of stomata: anisocytic and anomocytic. Panicoid type: three species showed this kind We also occasionally found paracytic stomata. This of epidermis: Digitaria sanguinalis, Echinochloa feature was also mentioned by Bruno et al. (1999), crusgalli, and Sorghum halepense. They were Cosa et al. (2000), Barboza et al. (2001), and characterized by rodlike micro-hairs. Digitaria Stenglein (2001). Solanum glaucophyllum presented presented dumb-bell silica-bodies while glabrous epidermis (Mansilla et al., 1999; Stenglein, Echinochloa nodular ones, and Sorghum dumb-bell, 2001), while the rest of species showed different nodular and cross shaped silica-bodies. Except for types of hairs, ie. (1) simple, conical 1-2-many celled the macro-hairs that we observed in Echinochloa in almost all the genera, (2) Y-shaped (in Physalis crusgalli, as Gould & Shaw (1992) did, the rest of and rarely in Salpichroa and Solanum), (3) dendritic features studied are completely in agreement with (in Physalis and rarely in Solanum), and (4) glan- Metcalfe´s results (1960). dular capitate (head 2-many celled) in almost all ge- nera. Our results agree with hair types found in Polygonaceae (7 toxic spp. analyzed): We found Datura by Carpano et al. (1990) and Licovsky et anomocytic stomata as the commonest type, but al. (2002). Our results accord to Metcalfe & Chalk there are also paracytic and anisocytic stomata. (1950, 1979) and Colares et al. (1999) who found Metcalfe & Chalk (1950, 1979) considered that the same hair types for Nicotiana, Physalis and paracytic stomata are not as frequent as anomocytic Salpichroa. Our results also agree with Cabrera but they do not mentioned anisocytic. We observed (1979) who found the same features in Physalis, papillae and different types of hairs: (1) simple, (2) and Cabrera (1979) and Stenglein (2001) in different conical-flagellate, (3) shaggy, (4) capitate, (5) species of Solanum. hydropoten, and (6) vesicular cells. Metcalfe & Chalk (1950) mentioned papillae, simple and On the basis of the stomatal types and size, uniseriate hairs and shaggy hairs (the last ones in anticlinal epidermal cell wall patterns, hair types and 1979). According to them, glandular hairs are wax ornamentation, the 180 species belonging to sessile. However, we found glandular hairs with two 41 families here surveyed can be differenciated by neck cells. Metcalfe & Chalk (1950) included within using the following key:
258 S. E. Freire et al., Epidermal characteristics of toxic plants for cattle Key to the toxic species of plants of Salado River Basin
1. Epidermal cells generally elongated. 1. MONOCOTYLEDONS
1'. Epidermal cells generally isodiametric. 2. DICOTYLEDONS
2. Leaves glabrous or papillose. Group A
2'. Leaves pilose. 3. Stomata paracytic. Group B
3'. Stomata not paracytic. 4. Leaves with glandular hairs exclusively. Group C
4'. Leaves with glandular and non-glandular hairs. 5. Glandular and non-glandular hairs in tufts (pilose nest) or hairs isolated, conical or whip. Group D
5'. Hairs isolated. Conical or whip hairs absent. 6. Bristles hairs present. Group E
6'. Bristles hairs absent. Group F
1. MONOCOTYLEDONS
1. Stomata anomocytic or tetracytic. 2. Stomata tetracytic. 1. Triglochin palustris
2'. Stomata anomocytic. 2. Habranthus tubispatus 3. Rhodophiala bifida
1'. Stomata paracytic. 3. Both macro- and micro-hairs present or at least one of them. 4. Macro- and micro-hairs present. 5. Micro-hairs with hemispherical distal cell. 6. Micro-hairs 14-20 µm long. Short cells solitary on intercostal zones. 4. Cynodon dactylon 5. C. hirsutus
6'. Micro-hairs 25-50 µm long. Short cells solitary or paired on intercostal zones. 6. Eragrostis cilianensis
259 Bol. Soc. Argent. Bot. 40 (3-4) 2005 5'. Micro-hairs with distal cell rod-like shaped. 7. Subsidiary cells domed. Papillae absent. 7. Digitaria sanguinalis
7'. Subsidiary cells triangular to dome. Papillae present. 8. Echinochloa crusgalli
4'. Micro-hairs or macro-hairs exclusively. 8. Micro-hairs exclusively. 9. Micro-hairs rod-like shaped. 9. Sorghum halepense
9'. Micro-hairs pear-like shaped or with hemispherical distal cell. 10. Short cells paired or in rows on costal zones. Papillae present. Subsidiary cells triangular to domed. Micro-hairs with hemispherical distal cell. 10. Leptochloa chloridiformis
10'. Short cells mostly solitary on costal zones. Papillae absent. Subsidiary cells trian- gular. Micro-hairs with distal cell pear-like shaped 11. Eleusine indica
8'. Macro-hairs exclusively. 11. Subsidiary cells parallel-sided. 12. Prickles absent. 12. Holcus lanatus
12'. Prickles present. 13. Hordeum murinum subsp. murinum
11'. Subsidiary cells domed. 14. Cortaderia selloana
3'. Macro-hairs and micro-hairs absent. 13. Prickles and hooks present. 14. With short cells on intercostal zones. 15. Elymus breviaristatus
14'. Without short cells on intercostal zones. 16. Lolium temulentum
13'. Prickles and hooks absent or prickles exclusively. 15. Prickles present. 16. Subsidiary cells parallel-sided. 17. Briza minor
16'. Subsidiary cells parallel-sided to dome. 18. Lolium multiflorum
15'. Prickles absent.
260 S. E. Freire et al., Epidermal characteristics of toxic plants for cattle 17. Subsidiary cells domed. 19. Cyperus rotundus
17'. Subsidiary cells parallel-sided to domed. 20. Lolium perenne
2. DICOTYLEDONS Group A: Leaves glabrous or papillose.
1. Stomata hexacytic. 1. Opuntia arechavaletae
1'. Stomata not hexacytic. 2. Stomata anomocytic. 3. Papillae on the veins present. 2. Ammi visnaga
3'. Papillae on the veins absent. 4. Licópoli glands present. 3. Limonium brasiliense
4'. Licópoli glands absent. 5. Hidropoten present. 4. Nymphoides indica
5'. Hidropoten absent. 6. Margin dentate. 7. Adaxial surface with cell type 3. 5. Polycarpon tetraphyllum
7'. Adaxial surface with cell type 1. 8. Both surfaces with cell type 1. Stomata more than 35 µm long. 6. Senecio bonariensis
8'. Abaxial surface with cell type 3. Stomata shorter than 35 µm long. 7. Senecio madagascariensis
6'. Margins not dentate. 9. Adaxial and abaxial surfaces with identical cell type. 10. Cell type 3/3. 11. Cuticular ornamentation present. 8. Conium maculatum
11'. Cuticular ornamentation absent. 12. Stomata shorter than 30 µm long. 9. Euphorbia peplus
12'. Stomata equal or longer than 30 µm long.
261 Bol. Soc. Argent. Bot. 40 (3-4) 2005 10. Cichorium intybus
10'. Cell type 1/1 or 2/2. 13. Cell type 2/2 11. Taraxacum officinale
13'. Cell type 1/1 14. Stomata shorter than 30 µm long. 12. Sonchus oleraceus
14'. Stomata longer than 30 µm long. 13. Phytolacca dioica
14. P. tetramera
9'. Adaxial and abaxial surfaces with different cell types. 15. Abaxial surface with only one cell type, 2 or 3. 16. Abaxial cell type 2. 17. Cuticular ornamentation present. Stomata 27-36 x 27-30 µm 15. Ranunculus cymbalaria
17'. Cuticular ornamentation absent. Stomata 20-30 x 20-25 µm 16. Passiflora caerulea
16'. Abaxial cell type 3. 18. Stomata 25-30 µm long. 17. Fumaria capreolata
18'. Stomata 30-45 µm long. 18. Ranunculus apiifolius
15'. Abaxial surface with cell type 1-2. 19. Adaxial cell type 1. Stomata 22-30 x 14-24 µm. 19. Euphorbia lathyrus
19'. Adaxial cell type 1-2. Stomata 25-35 x 20-30 µm. 20 Fumaria officinalis
21. Fumaria parviflora
22. Holmbergia tweedii
2'. Stomata not anomocytic. 22. Stomata paracytic, parallelocytic or diacytic (sometimes combined with stomata anomocytic). 23. Stomata paracytic or parallelocytic. 24. Stomata parallel to veins. 23. Eryngium paniculatum
24'. Stomata non-parallel to veins. 25. Stomata parallelocytic. Epidermal cells papillose.
262 S. E. Freire et al., Epidermal characteristics of toxic plants for cattle 24. Portulaca oleracea
25'. Stomata paracytic. Epidermal cells not papillose. 26. Stomata 35-45 x 20-30 µm. Bristles. 25. Galium richardianum
26'. Stomata 24-30 x 12-16 µm. 26. Manihot grahamii
23'. Stomata diacytic (sometimes combined with stomata anomocytic). 27. Papillae on veins present. 27. Ammi majus
27'. Papillae on veins absent. 28. Stomata diacytic exclusively. 28. Cyclospermum lept ophyllum
28'. Stomata diacytic and anomocytic. 29. Foeniculum vulgare
22'. Stomata anisocytic (sometimes combined with stomata anomocytic). 29. Stomata anisocytic exclusively. 30. Polygonum aviculare
29'. Stomata anisocytic and anomocytic. 30. Papillae present. 31. Papillae on the veins. 31. Rumex pulcher
31'. Papillae on all the surface. 32. R. obtusifolius
30'. Papillae absent. 32. Stomata longer than 40 µm. 33. R. crispus
32'. Stomata shorter than 40 µm. 33. Stomata shorter than 30 µm. Abaxial cell type 2-3. 34. Nicotiana glauca
33'. Stomata 30-35 µm long. Abaxial cell type 1. 35. Solanum glaucophyllum
Group B: Leaves pilose with stomata paracytic
1. Conical hairs. 2. Hairs ornamented. 3. Apical cell ornamented. 36. Oxypetalum solanoides
263 Bol. Soc. Argent. Bot. 40 (3-4) 2005 3'. All cells ornamented. 4. Abaxial surface with cell type 2. 37. Asclepias curassavica
4'. Both surfaces with cell type 1. 5. Cuticular ornamentation present. 38. A. mellodora
5'. Cuticular ornamentation absent 39. Morrenia brachystephana 40. M. odorata
2'. Hairs not ornamented. 6. Short conical hairs present. 41. Ricinus communis
6'. Short conical hairs absent. 7. Capitate glandular hairs present. 42. Convolvulus arvensis
7'. Capitate glandular hairs absent. 8. Stomata 20-25 µm long. 43 Salsola kali
8'. Stomata ca. 35 µm long. 44. Vinca major
1'. Other type of hairs. 9. Barrel shaped hairs. 45. Borreria verticillata
9'. Bristle hairs. 46. Senna corymbosa
Group C: Leaves with glandular hairs exclusively.
1. Glandular hairs not capitate. 2. Glandular hairs 1-seriate. 47. Trifolium repens f. repens
2'. Glandular hairs 2-seriate. 3. Glandular hairs sunken. 48. Hymenoxys anthemoides
3'. Glandular hairs not sunken. 49. H. cabrerae
1'. Glandular hairs capitate. 4. Head 1-celled. 5. Vesicular trichome (head prominent).
264 S. E. Freire et al., Epidermal characteristics of toxic plants for cattle 50. Chenopodium album 51. C. hircinum
5'. Head slightly developed. 52. Boerhavia diffusa var. leicarpa
4'. Head 1-2-many celled. 6. Glandular trichome with head many-radiate-celled. 53. Ibicella lutea
6'. Glandular trichome with head 1-2-celled. 7. Stomata anomocytic exclusively. 54. Anagallis arvensis
7'. Stomata anomocytic and anisocytic or polocytic 8. Stomata predominantly polocytic. 55. Plantago australis
8'. Stomata anomocytic and anisocytic. 56. Mirabilis jalapa
Group D: Leaves with pilose nest or with isolated hairs conical or whip.
1. Non glandular hairs associated with glandular hairs, forming pilose nest. 2. Ciclocytic stomata 3. Non glandular hairs with apical cell triangular. 57. Baccharis articulata
58. B. notosergila
3'. Non glandular hairs with apical cell tail-like. 59. B. rufescens
2'. Stomata anomocytic. 60. B. trimera
1'. Pilose nest absent. 4. Whip hairs. 5. Glandular hairs absent. 6. Whip hairs and other types of trichones. 7. Whip hairs on leaf surface and shaggy hairs on margin. 61. Carduus acanthoides
7'. Whip hairs and conical on leaf surfaces. 62. Aster squamatus
6'. Whip hairs exclusively.
265 Bol. Soc. Argent. Bot. 40 (3-4) 2005 8. Whip hairs ca. 9-celled. 63. Senecio tweediei 64. S. vulgaris
8'. Whip hairs 3-5-celled. 65. S. brasiliensis var. tripartitus 66. S. grisebachii 67. Baccharis artemisioides
5'. Glandular hairs present. 10. Glandular hairs capitate. 11. Head 1-celled (vesicular trichome). 68. Chenopodium murale
11'. Head many-celled. 69. Polygonum lapathifolium
10'. Glandular hairs not capitate. 12. Shaggy hairs present. 70. Cirsium vulgare
12'. Shaggy hairs absent. 13. Whip hairs with apical cell straight at the base. 71. Cynara cardunculus
13'. Whip hairs with apical cell bulbose at the base. 14. Whip hairs 2-4-celled. 72. Achyrocline satureioides 73. Centaurea solstitialis 74. Onopordon acanthium
14'. Whip hairs 3-11-celled. 75. Arctium minus 76. Centaurea calcitrapa 77. C. melitensis
4'. Conical hairs. 15. Vesicular cells present. 78. Polygonum convolvulus
15'. Vesicular cells absent. 16. Glandular hairs present. 17. Glandular hairs capitate. 18. Head 1-celled. 19. Glandular hairs with long stalk (ca. 6-celled). 79. Heliotropium amplexicaule
19'. Glandular hairs with short stalk (less than 6-celled).
266 S. E. Freire et al., Epidermal characteristics of toxic plants for cattle 20. Short conical hairs 1-celled. 80. Spergula arvensis
20'. Conical hairs 2-many celled. 21. Conical hairs 2-celled. 81. Geranium molle
21'. Conical hairs 2-many celled. 82. Solanum pygmaeum
18'. Head 2-many-celled (occasionally 1-celled mixed with many-celled). 22. Conical hairs 1-2-celled. 23. Stomata diacytic. 83. Lamium amplexicaule
23'. Stomata anomocytic and anisocytic. 84. Malva sylvestris
22'. Conical hairs 2-many-celled. 24. Y-shaped hairs present. 25. Asymmetrical Y-shaped. 85. Solanum sublobatum
25'. Symmetrical Y-shaped. 86. Salpichroa origanifolia
24'. Y-shaped hairs absent. 26. Conical hairs not ornamented. 87. Cestrum parqui
26'. Conical hairs ornamented. 27. Adaxial surface with cell type 1. 88. Datura ferox
27'. Adaxial surface with cell types 2 or 2 and 3. 89. Solanum chacoense 90. S. commersonii
17'. Glandular hairs not capitate. 28. Conical hairs ornamented. 29. Both surfaces with cell type 3. 91. Xanthium spinosum
29'. Both surfaces with cell types 1 and 2 or 2. 92. X. cavanillesii 93. Wedelia glauca
28'. Conical hairs not ornamented.
267 Bol. Soc. Argent. Bot. 40 (3-4) 2005 30. Cell types 1 and 2. 94. Ambrosia tenuifolia
30'. Cell type 3. 95. Bidens pilosa
16'. Glandular hairs absent. 31. Stomata anisocytic (sometimes combined with stomata anomocytic). 32. Conical hairs exclusively. 33. Conical hairs ornamented. 34. Hairs 1-celled. 35. Hairs longer than 400 µm long. 96. Sisymbrium irio
35'. Hairs shorter than 400 µm long. 97. Brassica rapa
34'. Hairs 2-3-celled. 36. Stomata anisocytic exclusively. Stomata 25-30 x 20-30 µm. 98. Adesmia bicolor
36'. Stomata anisocytic and anomocytic. Stomata 20-25 x 15-20 µm. 99. Melilotus officinalis
33'. Conical hairs not ornamented. 37. Hairs 1-celled. 38. Hairs on the surfaces. 39. Hairs shorter than 640 µm long. 100. Brassica nigra
39'. Hairs longer than 640 µm long (frequently 740-1300µm). 101. Sisymbrium altissimum
38'. Hairs on the margin exclusively. 102. Rorippa nasturtium-aquaticum
37'. Hairs 2-3-celled. 40. Conical hairs 3-celled. 103. Nicotiana longiflora
40'. Conical hairs 2-celled. 104. Lupinus gibertianus
32'. Conical and other types of hairs (non-glandular). 41. Hairs armed. 42. Stellate and 3-armed hairs. 105. Capsella bursa-pastoris
42'. Y-shaped and dendritic hairs.
268 S. E. Freire et al., Epidermal characteristics of toxic plants for cattle 106. Solanum diflorum
41'. Hairs not armed. 43. Barrel shaped. 107. Raphanus raphanistrum 108. R. sativus
43'. Falcate. 109. Cardaria draba
31'. Stomata anomocytic exclusively. 44. Conical hairs ornamented. 45. Hairs 1-celled. 110. Parietaria officinalis
45'. Hairs 3-6-celled. 46. All trichome cells ornamented. 111. Conyza bonariensis
46'. Apical cell ornamented exclusively. 112. Verbesina encelioides
44'. Conical hairs not ornamented. 47. Cuticula ornamented around the stomata present. 113. Gaillardia megapotamica
47'. Cuticula ornamented around the stomata absent. 48. Hairs 1-3-celled. 49. Apical cell bulbose at the base. 50. 1-celled hairs. 114. Echium plantagineum
50'. 3-celled hairs. 115. Bassia scoparia
49'. Apical cell not bulbose at the base. 116. Phytolacca americana
48'. Hairs 3-many-celled. 51. Papillae on the midvein present. 117. Amaranthus viridis
51'. Papillae on the midvein absent. 52. Both surfaces with cell type 1. 53. Apical cell with thick walls. 118. Solidago chilensis
53'. Apical cell with thin walls. 119. B. coridifolia
52'. Both surfaces with cell types 3 or with cell type 1 on adaxial surface, and type 3 on abaxial surface.
269 Bol. Soc. Argent. Bot. 40 (3-4) 2005 54. Adaxial surface with cell types 1, abaxial surface with cell types 3. 120. Amaranthus hybridus
54'. Both surfaces with cell types 3. 121. Tagetes minuta
Group E: Leaves with bristles hairs
1. Stomata diacytic. 122. Agrostemma githago
1'. Stomata not diacytic. 2. Peltate scales present. 123. Dodonaea viscosa
2'. Peltate scales absent. 3. Stinging hairs and cystholits present. 124. Urtica urens
3'. Stinging hairs and cystholits absent. 4. Glandular hairs present. 5. Glandular shaggy hairs present. 125. Turnera sidoides subsp. pinnatifida
5'. Glandular shaggy hairs absent. 6. Glandular clavate hairs 1-celled. 7. Cell type 1. 126. Clematis montevidensis
7'. Cell types 2 or/and 3. 127. C. bonariensis 128. Anemone decapetala
6'. Glandular capitate hairs. 8. Head 4-celled. 129. Melilotus albus
8'. Head 1-celled. 9. Bristles hairs ornamented. 10. Hairs 1-celled. 130. Oxalis corniculata
10'. Hairs 2-celled. 131. Melilotus indicus
9'. Bristles hairs not ornamented. 11. Hairs 1-celled. 12. Glandular capitate 1-celled with bottle shaped present. 132. Erodium malacoides
270 S. E. Freire et al., Epidermal characteristics of toxic plants for cattle 12'. Glandular capitate 1-celled with bottle shaped absent. 133. E. cicutarium
11'. Hairs 2-celled. 134. Vicia sativa subsp. nigra 135. Vicia graminea
4'. Glandular hairs absent. 13. Bristles hairs ornamented. 136. Parkinsonia aculeata
13'. Bristles hairs not ornamented (or only occasionally). 14. Bristles 1-celled. 15. Cuticular ornamentation present. 137. Sambucus australis
15'. Cuticular ornamentation absent. 16. Abaxial cell type 1. 138. Discaria americana
16'. Abaxial cell type 3. 139. Ranunculus repens var. repens 140. R. muricatus
14'. Bristles 2-celled. 141. Galega officinalis 142. Lotus corniculatus 143. L. glaber
Group F: Leaves with stellate, T-shaped, branched, shaggy, moniliform, or falcate hairs.
1. Glandular hairs present. 2. Glandular hairs not capitate. T-shaped hairs present. 3. Glandular hairs commonly present. Symmetrical T-shaped hairs. 144. Anthemis cotula
3'. Glandular hairs rarely present. Asymmetrical T-shaped hairs. 145. Vernonia rubricaulis
2'. Glandular hairs capitate. T-shaped hairs absent. 4. Glandular hairs with head 1-celled. 5. Falcate hairs, 1-celled. 146. Anchusa officinalis
5'. Stellate hairs, many-celled. 147. Solanum elaeagnifolium
4'. Glandular hairs with head 1-many-celled.
271 Bol. Soc. Argent. Bot. 40 (3-4) 2005 6. Stellate hairs present. 7. Porrect stellate hairs present. 148. Marrubium vulgare
7'. Porrect stellate hairs absent. 8. Stellate stalked hairs present. 149. Verbascum thapsus
8'. Stellate stalked hairs absent. 9. Stellate multiangulate hairs. Glandular hairs with head 2-celled. 150. Sida rhombifolia
9'. Stellate not multiangulate hairs. Glandular hairs with head 1-many-celled. 151. Solanum bonariense
6'. Stellate hairs absent. 10. Shaggy hairs. Vesicular cells and hydropoten present. 152. Polygonum persicaria
10'. Dendritic and Y-shaped hairs. Vesicular cells and hydropoten absent. 153. Physalis viscosa
1'. Glandular hairs absent. 11. Moniliform hairs present. 12. Stomata anomocytic. 154. Saponaria officinalis
12. Stomata parallelocytic. 13. Anticlinal cell wall very thick. 155. Wigginsia tephracantha
13'. Anticlinal cell wall thin. 156. Parodia ottonis
11'. Moniliform hairs absent. 13. Stomata anisocytic. Falcate hairs present. 157. Rapistrum rugosum
13'. Stomata anomocytic. Falcate hairs absent. 14. Shaggy hairs present. 15. Shaggy hairs on margin. Cell type 1. 158. Silybum marianum
15'. Shaggy hairs on margin and on midvein. Cell type 2. 159. Lactuca serriola
14'. Shaggy hairs absent (occasionally stellate hairs). 160. Fagara hyemalis
272 S. E. Freire et al., Epidermal characteristics of toxic plants for cattle Conclusions with an additional pair of lateral subsidiary cells) observed in the present study in Cactaceae. Hairs The following epidermal characters revealed of the families of this group are basically conical high diagnostic value for the identification of the (Amaranthaceae, Caryophyllaceae, toxic plants from the Salado River basin: 1) Chenopodiaceae, Phytolaccaceae) while the special hydropoten (Nymphoides indica); 2) licópoli glands type moniliform is present in Cactaceae and (Limonium brasiliense); 3) cystholits (Urtica Caryophyllaceae; 2) The close taxonomic urens); 4) parallelocytic stomata (Portulaca relationship between Convolvulaceae and oleracea, Wigginsia tephracantha, Parodia Solanaceae (order Solanales of Cronquist, 1981) is otonis); 5) polocytic stomata (Plantago australis); also supported by some of our data, e.g. conical 6) ciclocytic stomata (Baccharis spp.); 7) papillae hairs and capitate hairs, and stomata predominantly (Amaranthus viridis, Rumex spp.); 8) vesicular cells anisocytic and anomocytic; 3) Lamiaceae and (Polygonum spp.); 9) barrel shaped hairs (Borreria Scrophulariaceae (belonging to close related orders verticillata, Raphanus spp.); 10) vesicular hairs of Cronquist, 1981, Lamiales and Scrophulariales, (Chenopodium spp.); 11) glandular hairs with head respectively), have in common porrect hairs; 4) many-radiate-celled (Ibicella lutea); 12) hairs Geraniaceae and Oxalidaceae (order Geraniales of forming tufts (Baccharis spp.); shaggy hairs Cronquist, 1981) have bristles hairs; 5) Apocinaceae (Cirsium vulgare, Silybum marianum, Lactuca and Asclepiadaceae (order Gentianales of Cronquist, serriola, Polygonum lapathifolium); 13) Y-shaped 1981) share conical hairs and paracytic stomata. hairs (Solanum spp., Salpichroa origanifolia; 14) stellate hairs (Capsella bursa-pastoris, Verbascum Acknowledgements thapsus, Sida rhombifolia, Solanum spp); 15) peltate scales (Dodonaea viscosa); 16) stinging The authors appreciate the fine technical hairs (Urtica urens); T-shaped hairs (Anthemis assistance of Santiago M. Martínez, Vanesa G. cotula, Vernonia rubricaulis). Perrota and María B. Reitano (Facultad de Ciencias There are few characters which were not found Agrarias y Forestales, UNLP). The authors are also in the literature examined, and they could contribute grateful to the curators of herbaria LP and LPAG to improve the circumscription, from a anatomical for the permission to consult the specimens and point of view, the respective taxonomic group, i.e. María A. Migoya for inking our pencil original diacytic stomata in Apiaceae (Ammi, Cyclospermum, illustrations. We also thank to two anonymous Foeniculum); polocytic stomata in Plantaginaceae reviewers. Support for this study by the Consejo (Plantago); anisocytic stomata in Polygonaceae Nacional de Investigaciones Cientifícas y Técnicas (Polygonum, Rumex); and bristles in (CONICET), Argentina for S. E. F., E. U. and G. S., Caryophyllaceae (Agrostemma). and Programa de Incentivos, Secretaría de Políticas In addition, some epidermal characters analyzed Universitarias, Ministerio de Educación, is in this study, can be correlated with previously gratefully acknowledged. delimitated taxonomic groups above of family level within Dicotyledons: 1) Our epidermal characters Bibliography are consistent with the order Centrospermae (Caryophyllales order of Cronquist, 1981). In fact, AMAT, A. G. 1988. El uso de caracteres histofoliares en stomata are basically anomocytic in the relatively la identificación de las especies argentinas del género primitive families, (e.g. Amaranthaceae, Achyrocline DC. (Asteraceae). Acta Farm. Bonaeren- Chenopodiaceae, Nyctaginaceae, Phytolaccaceae). se 7: 75-83. Paracytic stomata, present in some species of ANCIBOR, E. 1984. Estudio anatómico de la vegetación Chenopodiaceae, can be related to the parallelocytic de la Puna de Jujuy y anatomía de Aschersoniodoxa mandoniana (Wedd.) Gilg. et. Muschler y Parodiodoxa stomata (i.e. paracytic stomata with an additional chionophila (Speg.) O. E. Schulz. Parodiana 3: 103- pair of subsidiary cells also parallel to the long axes 111. of the pore), of the more derivate families Cactaceae ANCIBOR, E. 1992. Anatomía ecológica de la vegetación and Portulacaceae. Stomata tetracytic present in de La Puna de Mendoza. I. Anatomía foliar. Parodiana Chenopodium spp. (Barboza et al., 2001) can be 7: 63-76. related to hexacytic stomata (i.e. tetracytic type
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276 S. E. Freire et al., Epidermal characteristics of toxic plants for cattle Appendix 1. Taxa and vouchers of toxic species used.
Amaranthaceae Amaranthus hybridus L. La Plata, Bayón 615 (LPAG). Amaranthus viridis L. La Plata, Bayón 603 (LPAG). Amaryllidaceae Habranthus tubispatus (L’Her.) Traub. Abra del Pantanoso Viejo, Pertusi 152 (LP). Rhodophiala bifida (Herb.) Traub. La Plata, Fabris 7039 (LP) Apiaceae Ammi majus L. Punta Lara, Cabrera 5710 (LP). Ammi visnaga (L.) Lam. Azul, Cabrera 9996 (LP). Conium maculatum L. Garín, Lanfranchi 211 (LP). Cyclospermum leptophyllum (Pers.) Sprague. Haedo, Spegazzini s. n. (LPS 20534 in LP). Eryngium paniculatum Cav. & Dombey ex F. D. Laroche. Sa. de la Ventana, Dawson et Núñez 148 (LP). Foeniculum vulgare Mill. La Plata, Cabrera 368 (LP). Apocinaceae Vinca major L. La Plata, Amorín in 1966 (LPAG).- Isla Martín García, Hurrell et al. 2374 (LP) - Laguna Brava, Cabrera et Fabris 17169 (LP). Asclepiadaceae Asclepias curassavica L. La Plata, Bayón 387 (LPAG). Asclepias mellodora A. St.-Hil. Gral. Villegas, Cabrera et Fabris 14789 (LP). Morrenia brachystephana Griseb. San Pedro, Fabris 3251 (LP). Morrenia odorata (Hook. & Arn.) Lindl. Pereyra, Cabrera 2058 (LP). Oxypetalum solanoides Hook. & Arn. Parque Pereyra Iraola, De la Torre in 1973 (LPAG). Asteraceae Achyrocline satureioides (Lam.) DC. San Clemente, Cabrera 4263, 4915 (LP). Ambrosia tenuifolia Spreng. Córdoba: San Javier, Bridarolli 1276 (LP) - Río Ceballos, Escalante 61 (LP). Anthemis cotula L. Pellegrini, Cabrera 6928 (LP) - Juancho, Cabrera 2744 (LP). Arctium minus (Hill.) Bernh. San Fernando, Lanfranchi 486 (LP) - La Plata, Cabrera 149 (LP). Aster squamatus (Spreng.) Hieron. var. squamatus. Buenos Aires. La Plata, Palo Blanco. Cabrera 138 (LP); Pellegrini, Cabrera 6984 (LP). Baccharis artemisioides Hook. & Arn.. Olavarría, Abbiatti 4095 (LP) - Pedro Luro, Cabrera 4514 (LP). Baccharis articulata (Lam.) Pers. Balcarce, Cabrera et Fabris 17.159 (LP) - Elizalde, Cabrera 1799 (LP). Baccharis coridifolia DC. Olavarría, Abbiatti 4015 (LP) - San Nicolás, Cabrera 7161 (LP). Baccharis. notosergila Griseb. Bavio, Zardini 593 (LP). Baccharis rufescens Spreng. Tandil, Hunziker 3912 (LP) - Salliqueló, Cabrera 7547 (LP). Baccharis trimera (Less.) DC. Magdalena, Zardini 592 (LP) - La Plata, Cabrera 464 (LP). Bidens pilosa L. var. pilosa. Delta del Paraná, Aº. Tuyuparé, Scala 427 (LP) – San Isidro, Cabrera 10647 (LP). Carduus acanthoides L. Monte Veloz, Cabrera 638 (LP) - Saladillo, Cabrera 6427 (LP). Centaurea calcitrapa L. Garín, Lanfranchi 576 (LP) - Sa. de la Ventana, Cabrera 5764 (LP). Centaurea melitensis L. Elizalde, Cabrera 534 (LP) – Magdalena, Cabrera 627 (LP). Centaurea solstitialis L. Gral. Conesa, Cabrera 4249 (LP). Cichorium intybus L. Gonnet, Delucchi 1757 (LP) – La Plata, Delucchi 1493 (LP). Cirsium vulgare (Savi) Ten. Pinamar, Cabrera 10078 (LP) - La Plata, Cabrera 140 (LP). Conyza bonariensis (L.) Cronsquist var. bonariensis. Otamendi, Hunziker 374 (LP). Cynara cardunculus L. La Plata, Cabrera 139 (LP).
277 Bol. Soc. Argent. Bot. 40 (3-4) 2005 Gaillardia megapotamica (Spreng.) Baker var. megapotamica. Río Negro: Cipolletti, Cabrera 703 (LP) – Neuquén: Portada Covunco, Cabrera 11090 (LP). Hymenoxys anthemoides (Juss.) Cass. Santa Fe: Laguna Paiva, Tur 647 (LP) – Leyes, Tur 455 (LP). Hymenoxys cabrerae K. L. Parker. Buenos Aires. Villarino, salitral de la Vidriera, Cabrera 6663 (LP). Lactuca serriola L. Prov. Corrientes. Santo Tomé, Krapovickas et al. 17033 (LP). Prov. Chaco. Cnia. Benítez, Schulz 178 (LP). Onopordon acanthium L. Magdalena, Cabrera 9159 (LP) – Entre Monte Veloz y Pipinas, Cabrera 622 (LP). Senecio bonariensis Hook. & Arn. Quilmes, Cabrera 319 (LP). Senecio brasiliensis (Spreng.) Less. var. tripartitus (DC.) Baker. Punta Lara, Dawson 867 (LP). Senecio grisebachii Baker var. grisebachii. Delta, Cabrera 4877 (LP). Senecio madagascariensis Poir. Mar del Plata, Solbrig 1073 (LP). Senecio tweediei Hook. & Arn. Punta Indio, Cabrera 24273 (LP) – Gral. Madariaga, Cabrera 10720 (LP). Senecio vulgaris L. Isla Maciel, Cabrera 938 (LP) - Miramar, Cabrera 5568 (LP). Silybum marianum (L.) Gaertn. Lincoln, Spegazzini 10341 (LP) - La Plata Cabrera 100 (LP). Solidago chilensis Meyen var. chilensis. Isla Martín García, Hurrell et al. 2875 (LP). Sonchus oleraceus L. San Clemente, Cabrera 4918 (LP). Tagetes minuta L. Tucumán: Tafí, Venturi 6111 (LP). Taraxacum officinale Weber ex F. H. Wigg. La Plata, Cabrera 176 (LP) – Isla Martín García, Hurrell 1934 (LP). Verbesina encelioides (Cav.) Benth. & Hook. f. La Plata, Cabrera 217 (LP) – Pinamar, Cabrera 10737 (LP). Vernonia rubricaulis Humb. & Bonpl. var. rubricaulis. Santa Fe: El Tostado, Job 1092 (LP). Wedelia glauca (Ortega) O. Hoffm. ex Hicken. Entre Ríos: S. Cerrito, Krapovickas et al. 22698 (LP). Xanthium cavanillesii Schouw. La Plata, Cabrera 1692, 175 (LP). Xanthium spinosum L. var. spinosum. Jujuy: La Quiaca, Cabrera et al. 15291 (LP) – El Carmen, Cabrera 7858 (LP). Boraginaceae Anchusa officinalis L. La Pampa: Santa Rosa, Fabris 1899 (LP) - Spegazzini s. n. (LPS 23884 in LP). Echium plantagineum L. Ensenada, Pérez Moreau in 1963 (LP) - La Plata, Monti 11 (LPAG). Heliotropium amplexicaule Vahl. Buenos Aires, Cabrera in 1943 (LP) - Gral. Alvarado, Fabris in 1960 (LP). Brassicaceae Brassica nigra (L.) W. D. J. Koch. Castelli, Alberto 6 (LPAG). Brassica rapa L. La Plata, Monti 15 (LPAG). Capsella bursa-pastoris (L.) Medik. Los Hornos, Bayón 588 (LPAG). Cardaria draba (L.) Desv. San Isidro, Cabrera 11565 (LP). Raphanus raphanistrum L. Otamendi, Hunziker 1488 (LP). Raphanus sativus L. Los Hornos, Bayón 589 (LPAG). Rapistrum rugosum (L.) All. La Plata, Monti 16 (LPAG). Roripa nasturtium-aquaticum (L.) Hayed. Tandil, Cabrera 6895 (LP). Sisymbrium altissimum L. Lomas sobre la margen izquierda del Río Limay, Chicchi 168 (LP). Sisymbrium irio L. Chubut: Isla de los Pájaros, Daciuk 63 (LP). Cactaceae Opuntia arechavaletae Speg. San Isidro (cultivado), Burkart 17911 (SI). Parodia ottonis (Lehm.) N. P. Taylor. Misiones, Teyucuaré, Burkart 15288 (SI). Wigginsia tephracantha (Link & Otto) D. M. Porter. URUGUAY: Dpto. Colonia, Estancia “Cerros de San Juan”, Kiesling 11593 (SI). Caprifoliaceae Sambucus australis Cham. & Schltdl. Buenos Aires, La Plata, Capelletti 85 (LPAG) - Bahía
278 S. E. Freire et al., Epidermal characteristics of toxic plants for cattle Samborombón, Ringuelet 206 (LPAG). Caryophyllaceae Agrostemma githago L. Agustina, Cabrera 6548 (LP). Polycarpon tetraphyllum (L.) L. Isla Martín García, Volponi 993 (LP). Saponaria officinalis L. Lincoln, Spegazzini in 1903. (LP). Spergula arvensis L. Entre Ríos: Colonia Adela, Bottino 463 (LP). Chenopodiaceae Bassia scoparia (L.) A. J. Scott. La Plata, Cabrera 7468 (LP). Chenopodium album L. Mar Chiquita, Dimitri in 1973 (LPAG). Chenopodium hircinum Schrad. var. hircinum. Punta Lara, Cabrera 4898 ( LP). Chenopodium murale L. La Plata, Bulta in 1972 (LPAG). Holmbergia tweedii (Moq.) Speg. Santa Fe: Santo Tomé, Job 1044 (LP). Salsola kali L. var. kali. San Clemente, Cabrera 4268 (LP). Convolvulaceae Convolvulus arvensis L. La Plata, Otero-Ríos in 1998 (LP). Cyperaceae Cyperus rotundus L. La Plata, Bayón 616 (LPAG). Euphorbiaceae Euphorbia lathyrus L. Boca del Riachuelo, Spegazzini s. n. (LPS 13929, 22794 in LP). Euphorbia peplus L. Isla Martín García, Hurrell 4117 (LP). Manihot grahamii Hook. Salta: Capital, Novara 2669 (LP). Ricinus communis L. Jujuy: Ledesma, Cabrera et Fabris 15985 (LP), Cabrera et Zardini 23840 (LP). Fabaceae Adesmia bicolor (Poir.) DC. Dolores, Arambarri 44 (LPAG). Galega officinalis L. Chascomús, Arambarri 197 (LPAG). Lotus corniculatus L. Buenos Aires. La Plata, Arambarri 61 (LPAG) - Entre Ríos: Concordia, Masut in 1972 (LPAG) . Lotus glaber Mill. Buenos Aires. La Plata, Arambarri 141 (LPAG) - Vieytes, Arambarri 220, Colares et al. s.n. (LPAG). Lupinus gibertianus C. P. Sm. Misiones: Cainguás, Leouncusat 948 (LP). Melilotus albus Desr. Castelli, Aguiar 2 (LPAG). Melilotus indicus (L.) All. Pilar, Dimitri in 1973 (LPAG). Melilotus officinalis (L.) Lam. Pilar, Arambarri in 1973 (LPAG) - Neuquén: Valle de Chasalnilla, Chicchi 100 (LP). Parkinsonia aculeata L. La Plata, Delucchi 1499 (LPAG). Senna corymbosa (Lam.) H. S. Irwin & Barneby. La Plata, Arambarri 242 (LPAG). Trifolium repens L. f. repens. La Plata, Delucchi 884 (LPAG). Vicia graminea Sm. var. graminea. La Plata, Pisano 96 (LP). Vicia sativa L. subsp. nigra (L.) Ehrh. La Plata, Arambarri et Perrotta 240 (LPAG). Fumariaceae Fumaria capreolata L. f. capreolata. La Plata, Monti 14 (LPAG). Fumaria officinalis L. Córdoba: Bajo Chico, Maldonado 101 (LP). Fumaria parviflora Lam. Carmen de Patagones, leg? (LP 21066). Geraniaceae Erodium cicutarium (L.) L’ Hér. ex Aiton. Chubut, Trevelín, Casaubón in 1979 (LPAG). Erodium malacoides (L.) L’ Hér. ex Aiton. Lavallol, Rinieri in 1966 (LPAG) - La Plata, Butta in 1972 (LPAG). Geranium molle L. La Plata, Arambarri 243 (LPAG). Juncaginaceae Triglochin palustris L. Isla Santiago, Spegazzini s. n. (LPS 16099 in LP). Lamiaceae
279 Bol. Soc. Argent. Bot. 40 (3-4) 2005 Lamium amplexicaule L. Vedia, Martínez 1 (LPAG) - La Plata, Amorín in 1966 (LPAG). Malvaceae Malva sylvestris L. Los Talas, Cabrera 1827 (LP). Marrubium vulgare L. Etcheverry, Martínez 2 (LPAG) – Dimitri in 1973 (LPAG). Sida rhombifolia L. Punta Lara, Cabrera 6342 (LP). Martyniaceae Ibicella lutea (Lindl.) Van Eselt. La Plata, Dimitri in 1972 (LPAG 7238). Menyanthaceae Nymphoides indica (L.) Kuntze. Tucumán, Santa Rosa, Venturi 616 (SI). Nictaginaceae Boerhavia diffusa L. var. leiocarpa (Heimerl) Adams. La Plata, leg? in 1988 (LPAG2349). Mirabilis jalapa L. Villa Elisa, Monti 12 (LPAG). Oxalidaceae Oxalis corniculata L. var. corniculata. La Plata, Arambarri 234 (LPAG) - La Plata, Cabrera 5353 (LP). Passifloraceae Passiflora caerulea L. La Plata, Vizcaíno in 1996 (LPAG) - Montes de Oca in 2001 (LPAG 5372). Phytolaccaceae Phytolacca americana L. Pereyra, Cabrera 2054 (LP) - Punta Lara, Landrum et Zardini 3052 (LP). Phytolacca dioica L. San Fernando, Isla Martín García, Hurrell et al. 2112 (LP) - La Plata, Pereyra in 1981 (LP). Phytolacca tetramera Hauman. Magdalena, Cabrera 1641 (LP). Plantaginaceae Plantago australis Lam. subsp. australis. Pta. Atalaya, Tur 1680 (LP). Plumbaginaceae Limonium brasiliense (Bois.) Kuntze var. brasiliense. Villarino, Cabrera 10171 (LP). Poaceae Briza minor L. La Plata, Bayón 568 (LPAG). Cortaderia selloana (Schult. & Schult. f.) Asch. & Graebn. La Plata, Bayón 605 (LPAG). Cynodon dactylon (L.) Pers. La Plata, Bayón 577 (LPAG). Cynodon hirsutus Stent. Luján, Nicora 629 (SI). Digitaria sanguinalis (L.) Scop. La Plata, Bayón 573, 574 (LPAG). Echinochloa crusgalli (L.) P. Beauv. La Plata, Bayón 581 (LPAG). Eleusine indica (L.) Gaertn. La Plata, Bayón 572 (LPAG). Elymus breviaristatus (Hitchc.) Á. Löve. Punta Indio, Rodrigo 3447 (LP). Eragrostis cilianensis (All.) Vignolo-Lutati ex Janch. La Plata, Bayón 576 (LPAG). Holcus lanatus L. Río Negro: Río Foyel, Bayón 590 (LPAG). Hordeum murinum L. subsp. murinum. El Talar, Lanfranchi 1 (SI). Leptochloa chloridiformis (Hack.) Parodi. Santa Fe: Logroño, Pire 1101 (SI). Lolium multiflorum Lam. La Plata, Bayón 592 (LPAG). Lolium perenne L. La Plata, Bayón 545 (LPAG). Lolium temulentum L. Entre Ríos: Concepción del Uruguay, Nicora 2023 (SI). Sorghum halepense (L.) Pers. La Plata, Bayón 575 (LPAG). Polygonaceae Polygonum aviculare L. Sa. de Olavarría, Abbiatti 4033 (LP) - Hurlingham, Schwabe 180 (LP). Polygonum convolvulus L. Garín, Lanfranchi 579 (LP) - Saliqueló, Cabrera 7510 (LP). Polygonum lapathifolium L. El Trigo, Cabrera 14725 (LP) - Sa. de la Ventana, Cabrera et Fabris 1 (LP). Polygonum persicaria L. BRASIL: Río Grande do Sul, Rambo 46510 (LPAG). Rumex crispus L. Isla Santiago, Cabrera 2222 (LP). Rumex obtusifolius L. Isla Martín García, Tur et. al 1810 (LP) - Sa. de Tandil, Delucchi 2198 (LP).
280 S. E. Freire et al., Epidermal characteristics of toxic plants for cattle Rumex pulcher L. Isla Martín García, Hurrell et al. 3877 (LP) - La Plata, Cabrera 5576 (LP). Portulacaceae Portulaca oleracea L. La Plata, Cabrera 527 (LP) - Isla Martín García, Hurrell et al. 3877 (LP). Primulaceae Anagallis arvensis L. San Nicolás, Cárdenas 7185 (LP) - Isla Martín García, Hurrell et al. 3874 (LP). Ranunculaceae Anemone decapetala Ard. var. decapetala. Tornquist, Proyecto Ventania 290 (LP) - Sa. de la Venta- na, Spegazzini in 1895 (LP). Clematis bonariensis Juss. ex DC. Punta Lara, Cabrera 2434 (LP). Clematis montevidensis Spreng. Sa. de las Tunas, Pertusi 60 (LP) - Corrientes: Bella Vista, Skorupka 10 (LP). Ranunculus apiifolius Pers. San Nicolás, Cabrera 7199 (LP)- Las Chilcas, Cabrera 8521 (LP). Ranunculus cymbalaria Pursh. Mar del Plata, Cabrera 9954 (LP). Ranunculus muricatus L. Isla Santiago, Cabrera 2263 (LP). Ranunculus repens L. var. repens. Atalaya, Tur 1531 (LP) - La Plata, Rodrigo 2814 (LP). Rhamnaceae Discaria americana Gillies & Hook. Tandil, Fabris et Schwabe 4738 (LP) - San Clemente, Cabrera 4928 (LP). Rubiaceae Borreria verticillata (L.) G. Mey. Lobería, Scala in 1918 (LP). Galium richardianum (Gillies ex Hook & Arn.) Endl. ex Walp. subsp. richardianum. Pellegrini, Cabre- ra 6934 (LP). Rutaceae Fagara hyemalis (Gillies) Engl. La Plata, Reitano 2002 (LPAG). Sapindaceae Dodonaea viscosa Jacq. Balcarce, Cabrera et Fabris 17.133 (LP). Scrophulariaceae Verbascum thapsus L. Tigre, Lanfranchi 430 (LP). Solanaceae Cestrum parqui L’ Hér. La Plata, Bayón 123 (LPAG). Datura ferox L. Benavídez, Lanfranchi 519 (LP). Nicotiana glauca Graham. La Plata, Bayón 371(b) (LPAG), Colares 6 (LPAG). Nicotiana longiflora Cav. La Plata, Bayón 381 (a) (LPAG). Physalis viscosa L. La Plata, Bayón 113 (LPAG). Solanum bonariense L. City Bell, Bayón 389 (a) (LPAG). Solanum chacoense Bitter subsp. chacoense. La Plata, Bayón 114 (LPAG). Solanum commersonii Dunal ex Poir. subsp. commersonii. Isla Martín García, Hurrell 3915 (LP). Solanum diflorum Vell. Vieytes, Bayón et Vizcaíno 466 (LPAG). Solanum elaeagnifolium Cav. La Plata, Butta in 1972 (LPAG) - Monte Hermoso, Ringuelet in 1942 (LPAG). Solanum glaucophyllum Desf. M. B. Gonnet, Ronco 1991 (LPAG) - Entre Los Porteños y City Bell, Bayón 175 (a) (LPAG). Solanum pygmaeum Cav. var. pygmaeum. Entre Ríos, Burkart 22721 (SI). Solanum sublobatum Willd. Vieytes, Bayón 468 (a) (LPAG). Salpichroa origanifolia (Lam.) Baill. City Bell, Bayón 139 (LPAG). Turneraceae Turnera sidoides L. subsp. pinnatifida (Juss. ex Poir) Arbo. Cnel. Suárez, Pertusi 76 (LP) - Sa. de la Ventana, Proyecto Ventana 899 (LP). Urticaceae Parietaria officinalis L. La Plata, Fabris in 1964 (LP). Urtica urens L. Lobería, Scala 1918. (LP).
281